Photoprotective acclimation of micro- and nano-size phytoplankton assemblages in the Indian sector of the Southern Ocean

Phytoplankton in the mixed layer is exposed to increasing levels of light when transported to the surface layer of the ocean. The photoprotective response of natural assemblages of phytoplankton can differ among community structures. We investigated photoprotective acclimation and xanthophyll cycle pigments in size-fractionated natural phytoplankton assemblages during the austral summer in the Indian sector of the Southern Ocean. We estimated concentrations of phytoplankton pigments in the micro-size fractions (>20 μm) and nano-size fractions (2–20 μm) by subtracting concentrations in the <20 μm fractions from concentrations in the bulk samples, and by subtracting concentrations in the <2 μm fractions from concentrations in the <20 μm fractions, respectively. Changes in the ratios of the xanthophyll cycle pigments diadinoxanthin (DD) and diatoxanthin (DT) were determined at three optical depths in the mixed layer and during 48 h deck incubations under solar photosynthetically available radiation and ultraviolet radiation. Large variations in (DD + DT)/Chl a in the mixed layer (percent coefficient of variation >67 %) and in deck incubation bottles under variable light conditions (>75 % of the temporal variation) for the micro-size fractions suggest a higher potential for photoprotective acclimation than for the nano-size fractions. Decreases in DT/(DD + DT) with increases in the optical depth of the mixed layer (ζ _MLD) suggest that larger variations in light availability in the mixed layer might predict lower values of DT/(DD + DT) at the surface, regardless of cell size.     

Temporal and spatial variation of the phytoplankton assemblage in the eastern Indian sector of the Southern Ocean in summer 2001/2002

The abundance and species composition of phytoplankton were investigated at stations in a permanently ice-free (61°S) and seasonally ice-covered area (64°S and 66°30′S) in the eastern Indian sector of the Southern Ocean between November 2001 and March 2002. Although a phytoplankton bloom occurred just after retreat of the sea ice at both stations in the seasonally ice-covered area, vertical stability of the water column during the bloom was weak at the most southerly station. This shows that a bloom can form even under weak vertical stability. In the bloom, diatoms dominated under weak vertical stability and Phaeocystis under strong vertical stability. In the latter case, ice algae largely contributed to development of the bloom. In the later observation period, a subsurface chlorophyll maximum (SCM) was observed at 61°S and 64°S. Species composition was different between the mixed layer and SCM at 64°S, but was uniform with depth at 61°S, indicating that the SCM is formed by different mechanisms.     

Size-fractionated primary production in the open Southern Ocean in austral spring

Size-fractionated primary production was measured by carbon-14 uptake incubations on three transects between 47°S and 59°30S along 6°W in October/November 1992. Open Antarctic Circumpolar Current and ice-covered Weddell Gyre water showed comparable low productivity (0.3 gCm^–2 day^–1) and size distribution. Picoplankton (<2 m) was the dominant size fraction, contributing approximately half to the total water column production. The significance of larger (>20 m) phytoplankton was only minor. Productivity in the Polar Front Zone north of 50°S, with higher water column stability, was up to 10 times higher with microplankton (>20 m) being predominant. No ice-edge bloom occurred over the 2 months study period; this is explained by non-favourable hydrographic conditions for blooming and the lack of melt-water lenses upon ice retreat. Picoplankton tended to make higher contributions at lower water column stability, and microplankton to make higher contributions at higher stability. Mixing, together with light climate, are discussed as the driving forces for Antarctic primary production and for its size structure.     

Size-fractionated Primary Production in the South Atlantic and Atlantic Sectors of the Southern Ocean

Results are presented from size fractionated chlorophyll a (Chla) and primary production studies along a transect between Antarcticaand southern Africa during the second South African AntarcticMarine Ecosystem Study (SAAMES II), conducted in late australsummer (January to February) 1993. Total integrated Chl a alongthe transect was highest in the vicinity of the Marginal IceZone (MIZ) and Antarctic Polar Front (APF). At these stations,integrated Chl a biomass was always >25 mg Chl a m^–2and was dominated by microphytoplankton. Although nominal increasesinChl a biomass were also associated with the Subantarctic Front(SAF) and Subtropical Convergence (STC), total Chl a biomassin these regions was dominated by nanophytoplankton. Withinthe inter-frontal regions, total integrated Chl a biomass waslower, generally <25 mg Chl a m^–2, and was always dominatedby nanophytoplankton. An exception was found in the AgulhasReturn Current (ARC) where picophytoplankton dominated. Totaldaily integrated production along the transect ranged between60 and 436 mg C m^–2 day^–1. Elevated production rateswere recorded at stations occupied in the vicinity of the MIZand at all the major oceanic frontal systems. The contributionsof the various size fractions to total daily production displayedthe same spatial pattern as integrated biomass, with microphytoplanktonbeing the most important contributor in areas characterizedby elevated phytoplankton biomass. Outside these regions, nanophytoplanktondominated the total phytoplankton production. Again, an exceptionwas found in the ARC north of the STC where picophytoplanktondominated total production. There, the lowest production alongthe entire transect was recorded, with total daily integratedproduction always <90 mg C m^–2 day^–1. The increasedproduction rates recorded in the MIZ appeared to result fromincreased water column stability as indicated by a shallow mixed-layerdepth. Within the inter-frontal regions, the existence of adeep mixed layer appeared to limit phytoplankton production.Low silicate concentrations in the waters north of the APF mayalso have limited the growth of large microphytoplankton.     

Seasonal variation of phytoplankton community structure and nitrogen uptake regime in the Indian Sector of the Southern Ocean

This study investigates the dynamics of phytoplankton communities and nitrogen uptake in the Indian sector of the Southern Ocean during spring and summer. The study area is oligotrophic (Chl a stocks <50 mg m⁻²); nevertheless, a large spatial variation of phytoplankton biomass and community structure was observed. During both seasons the phytoplankton community in the seasonal ice zone showed higher biomasses and was mainly composed of large diatom cells. However, in the permanently open ocean zone the community had low biomass and was chiefly composed of nano- and picoflagellates. In the polar front zone, although biomass was higher, the community structure was similar to the open ocean zone. The results suggest that the variation in phytoplankton community structure on a larger scale resonates with gradients in water column stability and nutrient distribution. However, significant changes in biomass and nutrient stocks but little change in community structure were observed. Absolute nitrogen uptake rates were generally low, but their seasonal variations were highly significant. During spring the communities displayed high specific nitrate uptake (mean rate = 0.0048 h⁻¹), and diatoms (in the seasonal ice zone) as well as nano- and picoflagellates (in the permanently open ocean zone and polar front zone) were mainly based on new production (mean ƒ-ratio = 0.69). The transition to summer was accompanied by a significant reduction in nitrate uptake rate (0.0048 h⁻¹ → 0.0011 h⁻¹) and a shift from predominantly new to regenerated production (ƒ-ratio 0.69 → 0.39). Ammonium played a major role in the seasonal dynamics of phytoplankton nutrition. The results emphasize that, despite a large contrast in community structure, the seasonal dynamics of the nitrogen uptake regime and phytoplankton community structure in all three subsystems were similar. Additionally, this study supports our previous conclusion that the seasonal shift in nitrogen uptake regime can occur with, as well as without, marked changes in community structure. Received: 2 December 1997 / Accepted: 20 April 1998     

Underwater light field and phytoplankton absorbance in different surface water masses of the Atlantic sector of the Southern Ocean

Spectral water transparency and phytoplankton light absorbance were studied in the Atlantic sector of the Southern Ocean during the Southern Ocean JGOFS ANT XIII/2 cruise in early austral summer 1995/1996. The study area comprised three zones, which differed markedly with respect to their hydrographic and planktological characteristics: the Antarctic Polar Frontal Zone with adiatom bloom, the Antarctic Circumpolar Current outside frontal systems with phytoplankton-poor water and a higher flagellate abundance than in the other two areas, and the marginal ice zone with a Phaeocystis bloom. The influence of phytoplankton on spectral water transparency was assessed by two independent procedures: the pigment-specific beam absorption coefficient, a_J*[5], at all stations, as estimated by spectroscopy of in vivo light absorption of plankton on glass fibre filters, and the pigment-specific light attenuation, (k_c[5]), as derived by regression analysis of spectral in situ vertical light attenuation coefficients in the sea against concomitant pigment concentrations. Values of a_J*[5] and vertical profiles of light attenuation by phytoplankton exhibited regional differences that corresponded with the three zones from which samples had been collected. These differences can be related to the specific characteristics of the three zones with respect to cell size distribution, pigment composition and biomass. The observed variations in a_J*[5] values should be considered when oceanic primary production is to be estimated by biooptical modelling.     

Natural age-dependent mortality rates of Antarctic krill Euphausia superba Dana in the Indian sector of the Southern Ocean

Data on the size and age composition of Antarctic krill (Euphausia superba Dana) were collected in the Cooperation and Cosmonaut Seas (Indian sector of the Southern Ocean) from 1985 to 1990. The estimation of the age-dependent annual extinction rate of krill [=1-exp(-M)] was obtained using the Zikov and Slepokurov (1982) approach and results were fitted by a parabolic equation. The coefficients of instantaneous natural mortality (M) of E. superba derived with this approach range from 0.52 during the maturation period, to 1.1–2.41 during the first and last years of life.     

Differences of the protozoan biomass and grazing during spring and summer in the Indian sector of the Southern Ocean

The dynamics of protozoa were investigated during two cruises in the Indian sector of the Southern Ocean: the early spring ANTARES 3 cruise (28 September to 8 November 1995) and the late summer ANTARES 2 cruise (6 February to 8 March 1994). Biomass and feeding activity of protozoa were measured as well as the biomass of their potential prey – bacteria and phototrophic flagellates – along the 62°E meridian. The sampling grid extended from the Polar Frontal region to the Coastal and Continental Shelf Zone in late summer and to the ice edge in spring, crossing the Antarctic Divergence. Protozoan biomass, although low in absolute terms, contributed 30% and 20% to the total microbial biomass (bacteria, phytoplankton and protozoa) in early spring and late summer, respectively. Nanoprotozoa dominated the total protozoan biomass. The geographical and seasonal distribution of protozoan biomass was correlated with that of phototrophic flagellates. However, bacterial and phototrophic flagellate biomass were inversely correlated. Phototrophic flagellates dominated in the Sea Ice Zone whereas bacteria were predominant at the end of summer in the Polar Frontal region and Coastal and Continental Shelf Zone. Furthermore, bacteria were the most important component of the microbial community (57% of the total microbial biomass) in late summer. Phototrophic flagellates were ingested by both nano-and microprotozoa. In contrast, bacteria were only ingested by nanoprotozoa. Protozoa controlled up to 90% of the daily bacterial production over the period examined. The spring daily protozoan ingestion controlled more than 100% of daily phototrophic flagellate production. This control was less strong at the end of summer when protozoan grazing controlled 42% of the daily phototrophic flagellate production. Accepted: 30 October 1999     

Spring phytoplankton assemblages in the Southern Ocean between Australia and Antarctica

Variations of phytoplankton assemblages were studied in November–December 2001, in surface waters of the Southern Ocean along a transect between the Sub-Antarctic Zone (SAZ) and the Seasonal Ice Zone (SIZ; 46.9°–64.9°S; 142°–143°E; CLIVAR-SR3 cruise). Two regions had characteristic but different phytoplankton assemblages. Nanoflagellates(<20 μm) and pico-plankton (∼2 μm) occurred in similar concentrations along the transect, but were dominant in the SAZ, Sub-Antarctic Front (SAF), Polar Front Zone (PFZ) and the Inter-Polar Front Zone (IPFZ), (46.9°–56.9°S). Along the entire transect their average cell numbers in the upper 70 m of water column, varied from 3 × 10⁵ to 1.1 × 10⁶ cells l⁻¹. Larger cells (>20 μm), diatoms and dinoflagellates, were more abundant in the Antarctic Zone-South (AZ-S) and the SIZ, (60.9°–64.9°S). In AZ-S and SIZ diatoms ranged between 2.7 × 10⁵ and 1.2 × 10⁶ cells l⁻¹, dinoflagellates from 3.1 × 10⁴ to 1.02 × 10⁵ cells l⁻¹. A diatom bloom was in progress in the AZ-S showing a peak of 1.8 × 10⁶ cells l⁻¹. Diatoms were dominated by Pseudo-nitzschia spp., Fragilariopsis spp., and Chaetoceros spp. Pseudo-nitzschia spp. outnumbered other diatoms in the AZ-S. Fragilaropsis spp. were most numerous in the SIZ. Dinoflagellates contained autotrophs (e.g. Prorocentrum) and heterotrophs (Gyrodinium/Gymnodinium, Protoperidinium). Diatoms and dinoflagellates contributed most to the cellular carbon: 11–25 and 17–124 μg C l⁻¹, respectively. Small cells dominated in the northern region characterized by the lowest N-uptake and new production of the transect. Larger diatom cells were prevalent in the southern area with higher values of N-uptake and new production. Diatom and nanoflagellate cellular carbon contents were highly correlated with one another, with primary production, and productivity related parameters. They contributed up to 75% to the total autotrophic C biomass. Diatom carbon content was significantly correlated to nitrate uptake and particle export, but not to ammonium uptake, while flagellate carbon was well correlated to ammonium uptake, but not to export. Diatoms have contributed highly to particle export along the latitudinal transect, while flagellates played a minor role in the export.     

Physiological state of phytoplankton communities in the Southwest Atlantic sector of the Southern Ocean, as measured by fast repetition rate fluorometry

The majority of the Southern Ocean is a high-nutrient low-chlorophyll (HNLC) ecosystem. Localized increases in chlorophyll concentration measured in the wake of bathymetric features near South Georgia demonstrate variations in the factors governing the HNLC condition. We explore the possibility that the contrast between these areas of high-chlorophyll and surrounding HNLC areas is associated with variations in phytoplankton photophysiology. Total dissolvable iron concentrations, phytoplankton photophysiology and community structure were investigated in late April 2003 on a transect along the North Scotia Ridge (53–54°S) between the Falkland Islands and South Georgia (58–33°W). Total dissolvable iron concentrations suggested a benthic source of iron near South Georgia. Bulk community measurements of dark-adapted photochemical quantum efficiency (F _v/F _m) exhibited a sharp increase to the east of 46°W coincident with a decrease in the functional absorption cross-section (σ_PSII). Phytoplankton populations east of 46°W thus displayed no physiological symptoms of iron or nitrate stress. Contrasting low F _v/F _m west of 46°W could not be explained by variations in the macronutrients nitrate and silicic acid and may be the result of taxon specific variability in photophysiology or iron stress. We hypothesize that increased F _v/F _m resulted from local relief from iron-stress near South Georgia, east of Aurora Bank, an area previously speculated to be a “pulse point” source of iron. Our measurements provide one of the first direct physiological confirmations that iron stress is alleviated in phytoplankton populations near South Georgia.     

Species-specific growth rate of phytoplankton in the West Pacific sector of the Southern Ocean. I.Prorocentrum scuttellum schröder

Phytoplankton samples were collected from the West Pacific Sector of the Southern Ocean to measure the growth rate from November 30 to December 1, 1995.Prorocentrum scuttellum was selected for growth rate measurement using the method of cell cycle analysis. During the 24 hr sampling cycle, cells ofP. scuttellum changed from 2,500 to 5,000 cells/L. The highest abundance was observed at 8:40 AM, December 1, and lowest at 11:40 PM, November 30. Cellular division seemed to occur sometime between 11:40 PM, November 30 and 2:40 AM, December 1. After cell division, DNA fluorescence shifted slowly towards the right, representing the S phase, and the majority of the cells were in S+G2 phases at 8:40 AM, December 1. Between the next six hours, a sharp drop in DNA fluorescence occurred, representing mitosis, and the majority of the cells returned to the G1 phase by 2:40 PM, December 1. We can not determine the duration time of the terminal event from this result However, the growth rate ofP. scuttellum was calculated as 0.43 d^-1 with the help of curve fitting methods. This unexpected result seems to have resulted due to background noise, unsynchronous cell division, unequal sampling, water column unstability, and migrating behavior ofP. scuttellum.     

Age,growth and reproductive biology ofNotothenia squamifrons gunther, 1880 from the Indian sector of the Southern Ocean

Summary The growth ofNotothenia squamifrons is analysed using scale reading of specimens from various isolated areas of the Indian sector of the Southern Ocean (Kerguelen, Crozet and Marion Islands; Ob, Lena and Kara-Dag seamounts). Studying reproductive biology reveals that spawning is annual and occurs during austral spring. Hatching takes place in summer. Scaling is set during the autumn. The smallest scaled fish measure 3.8 cm in total length. The data mentioned above, the check number on the scales and the sampling dates lead to the identification of the different age groups. Differences in growth rate appear between the studied areas. The fastest growth is observed around the Kerguelen Islands where the oldest specimens are 19 years old and reach 52 cm in total length. On the other hand, in the Crozet population,N. squamifrons does not live more than 12 years for a maximum length of 30 cm. Scale reading in fish from the Ob and Lena seamounts shows a minor decrease in growth during the winter season when compared to that in the other areas. The hydrographical conditions of this sector of the Southern Ocean may, in part, explain these differences: especially temperature and plankton biomass seem to effect growth rate. The cold antarctic waters and the highest plankton productivity, inducing the optimal growth rate, are found in the Kerguelen Islands.     

Summer distribution of netphytoplankton in the Atlantic sector of the Southern Ocean

The surface distribution of netphytoplankton (>20 m) in the Atlantic sector of the Southern Ocean was investigated along two transects during early and late austral summer 1990/91. Sampling was under-taken at intervals of 60 of latitude between 34° and 70°S for the analysis of nutrients and for the identification and enumeration of netphytoplankton. Peaks in total diatom abundances were recorded at the Antarctic Polar Front (APF), in the vicinity of the South Sand wich Islands, in the marginal ice zone and in the neritic waters of the Atlantic sector of Antarctica. Cluster analysis indicates the existence of two major zones between Southern Africa and Antarctica. Diatom abundance increased south of the Antarctic Polar From along both transects, which can be partially explained by gradients of silicate concentration. Small chain-forming species (e.g. Fragilariopsis kerguelensis and Nitzschia lineata) dominated the diatom assemblages in early summer, while larger species, such as Rhizosolenia hebetata f. semispina and Corethron criophilum, dominated late summer diatom assemblages. The predominance of typically ice-associated forms in early summer suggests that the release of epontic cells during ice melt provides the initial inoculum for the netphytoplankton biomass. These small diatoms are subsequently replaced by larger species.     

Deep-sea pelagic ichthyonekton diversity in the Atlantic Ocean and the adjacent sector of the Southern Ocean

Aim Deep‐sea pelagic diversity is poorly understood. Local (SL) and regional (SR) ichthyonekton species richness are presented and analysed with respect to local and regional environmental factors, and biogeographical processes. Location Sixty‐six stations from the Atlantic Ocean and adjacent sector of the Southern Ocean, 65° N to 57° S. Methods Estimation of SL by means of rarefaction. Stepwise evaluation of SL and SR relationships by means of the second‐order corrected Akaike information criterion (AICc) after locally weighted scatterplot smoothing (LOESS) and linear fitting, analysis of saturation effects by means of slopes of species accumulation curves (log–log plots). Results Latitudinal gradients were present for SL and SR, and were asymmetric between the Northern and Southern hemispheres. Relatively low species richness was encountered for the Southern Ocean. Asymmetry at the regional level by means of higher SR was attributed to area effects in the South Atlantic. Log–log plots indicated saturation of local assemblages and dependence on environmental factors. SL was related to productivity; this relationship was hump‐shaped. SR was positively related to area size and negatively to seasonality of production. Biogeographical effects were indicated in that SR peaks coincided with overlap zones of boreal and tropical faunas as a consequence of historical faunal exchange processes. Main conclusions The stepwise approach allowed for distinction between effects of area size, productivity and biogeographical processes on diversity at local and regional scales. Productivity in particular is important in two ways. At the local scale, the link of productivity to SL is explained by a successional‐functional hypothesis of resource utilization, whereas the seasonality effect for SR reinforces the hypothesis of dependence of deep‐sea fishes on seasonality of production through changes of life‐history traits. The causes of low Antarctic faunal diversity remained unresolved.     

Biology and population dynamics of Notothenia rossii rossii from the Kerguelen Islands (Indian sector of Southern Ocean)

Summary Recent cruises on board research vessels and trawlers have followed the adult fishes of the Notothenia rossii rossii Richardson, 1844 population from the Kerguelen Islands. The food of this Nototheniidae being mainly planktonic, the movements of shoals are connected with the high productivity areas. In the Southern Ocean these areas are localized around the Antarctic Convergence. At the Kerguelen Islands the irregular configuration of the shelf and seasonal variations in water masses change the positions of the productive areas. For this reason the adult population can be to the north or south off the archipelago. Spawning only gathers the mature fishes in the southeast of Kerguelen during winter. At that time this species is very vulnerable to fishing efforts. The population dynamics of this Antarctic fish is studied with fishery statistics and biological data. A decline of the average length and catches per unit effort are observed from one year to the next.

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Resumé De récentes missions à bord de navires de recherche et de chalutiers ont permis de suivre la population adulte de Notothenia rossii rossii Richardson, 1844 des îles Kerguelen. Le régime alimentaire de ce Nototheniidae étant principalement de nature planctonique, les mouvements des bancs de poissons sont liés aux zones de haute productivité. Dans l'océan austral ces zones sont localisées au niveau de la convergence antarctique. Aux îles Kerguelen la topographie du plateau continental et les variations saisonnières de l'hydrologie modifient continuellement la position des zones de productivité. Pour cette raison la population adulte peut se trouver soit au nord soit au sud de l'archipel. Seule la ponte réunit les poissons matures dans le sud-est des îles Kerguelen pendant l'hiver. A cette période l'espèce est très vulnérable à un effort de pêche. La dynamique de la population de ce poisson antarctique est étudiée à partir des statistiques de pêche et des données biologiques. Une baisse de la longueur moyenne de la population ainsi qu'une diminution des valeurs de la prise par unité d'effort sont observées pendant la période considérée.

     

Carbon demand,utilization, and metabolic diversity of bacterioplankton in the frontal regimes of the Indian sector of the Southern Ocean

Bacterial production, respiration and metabolic diversity were measured up to 120 m depth in the Sub-Antarctic Front (SAF) and Polar Fronts I and II (PFI and PFII) of the Indian Ocean sector of the Southern Ocean during 2010 Austral Summer. Prokaryotic cell count was maximum at PFI and PFII (~109 cells L−1) and minimum at SAF (~107 cells L−1). Furthermore, integrated bacterial production was higher at PFI (1.07 mg C m−2 h−1) and PFII (0.72 mg C m−2 h−1) compared to SAF (0.61 mg C m−2 h−1). At PFII, integrated bacterial growth efficiency was higher (8.96) compared to PFI (7.42) and SAF (7.17), signifying that the net contribution of PFII to the microbial loop could be relatively pronounced. Enhanced cell numbers and production at polar fronts indicate that the dissolved organic matter could be converted to secondary biomass through the microbial loop. However, integrated bacterial respiration rate at PFII (0.83 mg C m−2 h−1) was lower than that at PFI (1.84 mg C m−2 h−1) resulting in higher growth efficiency at PFII. Metabolic flexibility at SAF was clearly brought about by utilization of carboxylic acids like D-malic acid and itaconic acid, and carbohydrates like N-acetyl D-glucosamine, D-cellobiose and D-lactose. Utilization of amino acids like glycyl L-glutamic acid and L-threonine, and an amine, phenylethylamine, was critical in determining the metabolic variability at PFI. PFII hosted microbes that utilized phenolic compounds (2-hydroxy benzoic acid and 4-hydroxy benzoic acid) and polymers (like Tween 80). Utilization of polyols over carbohydrates in polar waters indicates a niche with lesser influence of the Antarctic melt waters on the bacterioplankton metabolism.